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Analysis of Ontogenetic Changes in Head Shape

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Analysis of Ontogenetic Changes in Head Shape Analysis of ontogenetic changes in head shape and diet in a catfish with moderately enlarged jaw adductors (Clariallabes melas) Marisa Wyckmans, Anthony Herrel, Dominique Adriaens To cite this version: Marisa Wyckmans, Anthony Herrel, Dominique Adriaens. Analysis of ontogenetic changes in head shape and diet in a catfish with moderately enlarged jaw adductors (Clariallabes melas). Belgian Journal of Zoology, Royal Belgian Zoological Society, 2011, 141 (2), pp.11-20. hal-02436104 HAL Id: hal-02436104 https://hal.archives-ouvertes.fr/hal-02436104 Submitted on 15 Jan 2020 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. Belg. J. Zool., 141 (2) : 11-20 July 2011 Analysis of ontogenetic changes in head shape and diet in a catfish with moderately enlarged jaw adductors (Clariallabes melas) Marisa Wyckmans1, Anthony Herrel2* & Dominique Adriaens3 1 Dept. Biology, University of Antwerp, Universiteitsplein 1, B-2610 Antwerpen, Belgium. E-mail: [email protected] 2 Dept. d’Ecologie et Gestion de la Biodiversité, Muséum National d’Histoire Naturelle, 57 rue Cuvier, 75231, Paris, France. E-mail: [email protected] 3 Dept. Biology, Ghent University, K.L. Ledeganckstraat 35, B-9000 Gent, Belgium. E-mail: [email protected] * Corresponding author: [email protected] ABSTRACT. Jaw adductor hypertrophy, or the presence of enlarged jaw-closing muscles, has arisen several times independently in African clariid catfish. Previous work has demonstrated that species characterized by enlarged jaw adductors may have unusual ways of foraging such as terrestrial foraging and prey capture, and often include a large proportion of hard and terrestrial prey in their diet. However, relatively little is known about species of the genus Clariallabes with an intermediate degree of jaw adductor hypertrophy. In the present study we present data on head shape and diet for a range of sizes of specimens of a poorly known species of Clariallabes, C. melas. Our data show that growth patterns in this species and the previously studied C. longicauda are similar in some ways (e.g. positive allometry of the growth of the jaw muscles) but different in others (negative allometry in hyoid width). However, C. melas has a smaller head for its body size in all dimensions. Due to the large number of empty stomachs we encountered, dietary data remain preliminary, but suggest a varied diet including both hard and soft prey. Our data show that a large amount of variation in head shape may exist even among related species and that species with jaw adductor hypertrophy generally show positive allometry in the growth of the jaw adductors and associated structures. Whether jaw muscle hypertrophy is an adaptive trait in clariid catfish awaits further comparative analyses testing for the evolutionary association between jaw adductor hypertrophy and the inclusion of hard prey into the diet. KEY WORDS: fish, muscle, biting, suction, feeding, ontogeny INTRODUCTION (LAUDER, 1983; KORFF & WAINWRIGHT, 2004). Interestingly, hypertrophy of the jaw adductors In fish, the inclusion of hard prey into the diet is often noted in populations showing a higher (durophagy) is often associated with changes proportion of molluscs in the diet, suggesting in dentition (e.g. STREELMAN & ALBERTSON, an adaptive evolutionary response of the cranial 2006), the presence of enlarged pharyngeal morphology to diet (TURINGAN et al., 1995). or oral jaw adductor muscles (WAINWRIGHT, Moreover, a strong plastic response in muscle 1987; HERNANDEZ & MOTTA, 1997), an size to an increase of molluscs in the diet has overall increase in the robustness of the oral been demonstrated in some fish M( ITTELBACH et or pharyngeal jaws and the skull (CABUY et al., 1999). al., 1999; GRUBICH, 2003) and in some species durophagy is associated with the presence Oral jaw adductor hypertrophy has arisen of a specialized muscle activation pattern several times independently in African clariid 12 Marisa Wyckmans, Anthony Herrel & Dominique Adriaens catfish (JANSEN et al., 2006; DEVAERE et al., WASSENBERGH et al., 2006a), and that they 2007). Indeed, a recent study using combined include a large proportion of hard and often morphological and molecular data suggests that terrestrial prey in their diet (HUYSENTRUYT an increase in jaw adductor size has originated at et al., 2004). However, the species with least four times independently (DEVAERE et al., the most extreme degree of jaw adductor 2007). Clariid catfish are, however, not known hypertrophy appear to swallow their prey to be mollusc crushers and retain large pointed whole (HUYSENTRUYT et al., 2004). Previous teeth on the jaws (DEVAERE et al., 2001). Among studies have shown that rapidly-growing and clariid catfish species, jaw adductor hypertrophy suction-feeding clariid species, such as Clarias ranges from the normal dorsoventrally- gariepinus grow isometrically (HERREL et al., compressed type with the jaw adductors largely 2005). In contrast, the growth of jaw adductors covered by the cranial bones (e.g. within the and related structures shows positive allometry genus Clarias), to a situation where most of the in a species with an intermediate degree of jaw dorso-lateral cranial bones are strongly reduced hypertrophy, such as Clariallabes longicauda to make space for the jaw adductor muscles (WYKMANS et al., 2007). In order to test the (e.g. within the genus Channallabes) (DEVAERE hypothesis that species with intermediate degrees et al., 2001). Although the species with more of jaw hypertrophy are characterised by positive pronounced degrees of jaw adductor hypertrophy allometric growth, we analysed growth patterns have been relatively well studied, little is known in a species with an intermediate degree of jaw about species with an intermediate degree of jaw adductor hypertrophy (Clariallabes melas; adductor hypertrophy, such as those of the genus BOULENGER 1887). Moreover we compare Clariallabes (but see WYCKMANS et al., 2007). patterns of growth in C. melas to those observed for its congener, C. longicauda (BOULENGER, In other vertebrates, an increase in the size of the 1902) and present data on diet for a range of oral jaw adductor muscles is typically associated sizes of individuals of C. melas. with an increase in bite force (HERREL et al., 2007a), and in many vertebrates an increase in bite force is associated with the inclusion of MATERIALS AND METHODS hard or large prey into the diet (HERREL et al., 2002; AGUIRRE et al., 2003; HERREL et al., 2005; Material examined HUBER et al., 2009). However, at least for some groups such as lizards, it has been shown that The 51 Clariallabes melas used in this study sexual selection for increased fighting capacity were obtained from the Royal Museum for may also lead to increased bite force (LAILVAUX Central Africa (RMCA - KMMA), Tervuren, et al., 2004; HUYGHE et al., 2005; LAPPIN & Belgium. All specimens (30592-603-1 to 10; HUSAK, 2005) and sexual differences in the size 68800-68802-1 to 2; 117776-117780-1 to 5; of the jaw adductors (HERREL et al., 2007b). 119100-101-1 to 2; 29660; 46746-46747-1 to Moreover, in some burrowing mammals, high 2; 117770-117775-1 to 2; 178220-230-MEL15, bite force may be associated with the ability to 178220-230-1; 19072-19074-1; 38495-38508-1 dig tunnels into hard soils rather than diet (VAN to 2; 44916-917-1 to 2; 74381; 79220; 96646- DAELE et al., 2009) suggesting that a multitude 96648-1 to 2; 98931; 119117; 138665; 73-68-P- of selective pressures may be operating on jaw 566-574-1 to 8; 86-06-P-51-53-1 to 2; 88-01-P- muscle size and bite force capacity. 1993; 88-01-P-1994-1995-1 to 2) were collected in the Congo River drainage. Fish included in Previous work on clariid catfish with enlarged this study range from 116 to 329mm total length oral jaw adductors has demonstrated that and encompass juvenile (i.e. non-reproductive) some of these species have unusual ways of and adult specimens. foraging including terrestrial prey capture (VAN Head shape and diet in Clarriallabes 13 Morphometrics greater degree, they should be more successful in capturing elusive prey. Body size (both total length and standard length) and head dimensions were measured Diet using digital calipers (Mitutoyo CD-30C and CD- 15B; ± 0.01mm). Head length was measured as Stomach contents were removed in situ through the distance between the tip of the snout and the a ventral incision, and preserved in a 70% aqueous caudal edge of the occipital process. Head width ethanol solution. All prey items were sorted and and head height were measured just posterior to identified using a binocular microscope (Wild the jaw adductors. M3Z). Because the majority of organisms found in the stomachs were crushed and/or digested, The surface area taken up by the jaw adductors they were identified to the level of order or higher in dorsal view was determined using digital taxonomic level where appropriate. photographs of each specimen and is used here as a proxy for jaw adductor size (see Fig. 1; The number of prey items found in every WYCKMANS et al., 2007). For our analyses, the stomach was counted and for every prey item average of the left and right sides was calculated. prey size was estimated. Intact prey items were Additionally the maximal width of the head and measured using digital calipers (Mitutoyo CD- the width of the neurocranium were determined 15B; ± 0.01mm).
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